Predators and Parasites of the Douglas-Fir Beetle: Description and Identification of the Immature Stages

TECHNICAL BULLETIN 79 OCTOBER 1964

Predators and Parasites of the Douglas-fir Beetle: Description and Identification of the Immature Stages

DEC 1964 LIBRARY ION STATE

NNiVOSTV .

Agricultural Experiment Station Oregon State University Corvallis CONTENTS

Page INTRODUCTION 3

LITERATURE REVIEW . . 4 IDENTIFICATION OF THE INSECT ENEMIES OF DENDROCTONUS PSEUDOTSUGAE . 5 Methods and Procedures of Study 5

Known and Possible Predators and Parasites . 7 Field Keyto the Insect Larvae of the Known Predators and Parasites Description and Illustration of the Egg,Larva,and Pupa

of the Known Predators and Parasites . 9

Cleridae: Enoclcrus sphegeus Fabricus . 9

Cleridae: Enoclcrus lecontei Wolcott . 19 Cleridae:Thanasimus undatulus Say 25

Ostomatidae:Temnochila virescens chlorodia Mannerheim 32 Dolichopodidae:NVcdetcra spp 37

Lonchaeidae: Lonchaca spp 44

Braconidae:Coeloides brunneri Viereck . 46

Pteromalidae:Roptrocerus eccoptogasteri Ratz, Cecidostiba burkciCraw.,and C dendroctoni Ash 49

ACKNOWLEDGMENTS . 50

REFERENCES CITED. . 50

AUTHORS :L. N. Kline is a former research assistantand J. A. Rudinsky is a professor of forest entomology, Department of Ento- mology, Oregon StateUniversity.This bulletin is based on the senior author's Master's thesis submitted to OSU. Predators and Parasites of the Douglas-fir Beetle:

Description and Identification of the Immature Stages L. N. KLINE AND J. A. RUDINSKY

INTRODUCTION The Douglas-fir beetle,Dendroctonus pseudotsugaeHopkins (Coleoptera: Scolytidae), is the most destructive insect enemy of Douglas-fir,Pseudotsuga menziesii(Mirb.) Franco, throughout the range of this important tree species. During the epidemic of 1951- 1954, for instance, it killed over three billion board feet of green timber in western Oregon and Washington alone. The beetle is native to North America, and is preyed upon by a number of important predators and parasites. Some of these undoubtedly play a significant role in reducing the epidemic population of the beetle or in preventing its increase to destructive numbers. Making a prognosis of the trend of the Douglas-fir beetle popu- lationin the field or forecasting the economic damage requires sampling not only Douglas-fir beetles, but also numbers and species of the immature stages of predators and parasites present under the bark. Past studies have been concerned more with the biology of the Douglas-fir beetle than with its natural enemies. The biology and identification, especially of the immature stages of the predators and parasites, are incomplete or in some instances virtually unknown. This bulletin presents identifications, descriptions, and drawings of the immature stages of the main predators and parasites of the Douglas-fir beetle in the Intermountain and Pacific Northwest regions of the United States. Brief notes on the life histories of the predators and parasites, and a key separating the larvae are also included. Emphasis is placed on the following species :Enoclerus sphegeus, E. lecontei, Thanasimus undatulus,andTemnochila virescens.The remaining species,Medeteraspp.,Lonchaeaspp.,Coeloides brunneri, and the pteromalids, are treated in a more general manner. Three instars exist for the clerids, with five to six for the ostomatid. LITERATURE REVIEW A review of the literature shows that only limited studies have been made of the biology, habits, and identification of a few of the insect enemies of the Douglas-fir beetle. In northern Idaho and Montana, Bedard (1, pp. 27-61) worked briefly on the biology, habits, and identification ofEnoclerus sphegeus Fab. (Coleoptera :Cleridae) ;E. leconteiWolc. (Coleoptera :Cleri- dae) ;T hanasivnus dubius(Fab.) (Coleoptera: Cleridae), a mistaken identity forT undatulus Say; Teznnochila virescens chlorodiaMann. (Coleoptera :Ostomatidae) ;Medetera aldrichiiWh. (Diptera :Doli- chopodidae);Lonchaea corticisTay. (Diptera: Lonchaeidae);Coe- loides brunneriVier. ('Hymenoptera; Braconidae);Roptrocerus ec- co ptogasteriRatz. (Hymenoptera :Pteromalidae) ;andCecidostiba dendroctoniAshm. (Hymenoptera: Pteromalidae). Bedard'sde- scriptions and drawings of the larvae were only of the mature instars. Roving, Craighead, and Champlain(4, 5, 6)published on the identification of the mature larval instars of E.sphegeus, E. lecontei, andT. virescens.Only a few drawings were made forT. virescens. These workers also included brief notes on the biology of some of the clerids preying on bark beetles in general. Habits of the adults and larvae and identification of four instars of E.sphegeusin relation toIps pini(Say) andIps perrottiSw. on lodgepole pine slash in Alberta, Canada, have been reported by Reid (20). Person(17) worked on the life history, habits, importance as a natural control factor, and possibilities of increasing the effectiveness of E.lecontei in association with western pine beetle,Dendroctonus brevicosnis Lec., in California. Some work has been done by Struble and Car- pelan on the identification, biology, habits, and laboratory propagation of E.sphegeusandT. virescensin an attempt to rear these predators for liberation in the field as control agents of bark beetles (24, 25, 26). A fairly complete study was made by DeLeon (8) on the biology, habits, and internal and external morphology of the larva ofMedetera aldrichiiWh. in relation to mountain pine beetle,D. nionticolaeHopk., in Montana and northeastern Washington. Also, several notes were made by Hopping (9) on the seasonal development ofM. aldrichiias a predator ofD pseudotsugaein British Columbia. Recently, Johnsey(10)studied the biology ofM. aldrichiiand Lonchaeasp. associated with the Douglas-fir beetle in westernOregon and Washington. Ryan and Rudinsky (22) published on the biology, habits, and descriptions of the five larval instars, development of the immature stadia at various temperatures, and factors contributing to the effectiveness ofCoeloides brunneriVier., a parasite of the Douglas-fir beetle in western Oregon.

IDENTIFICATION OF THE INSECT ENEMIES OF DENDROCTONUS PSEUDOTSUGAE As just noted, previous studies, with the exception of those by DeLeon (8, pp. 62-67), Reid (20), and Ryan (21), were concerned only with the identification of the mature larval stage. Past and present biological appraisals or population sampling of bark beetles have been based on immature stages found under the bark. There- fore, in this type of sampling, since all stages of the insect concerned may be encountered, all should be easily identified. Discussed here are the methods and procedures used in rearing certain species in order to obtain specimens for study, and also the methods of measurement and drawing. To facilitate future identifications, a field key, descriptions, and illustrations of the immature stages of the most important species are included. The general biology, geographical distribution, and relative importance of each species are given, based on general observation.

Methods and Procedures of Study Because of the scant taxonomical knowledge of the adults of the hymenopterous parasites and dipterous predators, it was possible in this study to identify the immature stages only to the family and generic levels. Available literature was used for the descriptions whenever possible, and other characters and material were included from personal observation. The taxonomy of the adult coleopterous predators has been worked on more intensively but still presents a problem in associating adults with their immature forms. Therefore, each species was reared in order to obtain known specimens of each stage for study. These rearings were conducted in a manner similar to those reared by Struble (25). Briefly, the method of rearing was as follows : Known identified adults ofEnoclerus sphegeus, E lecontei, Thanasirnus undatulus,and Temnochila virescenswere collected in the field on the bark and around Douglas-fir trees infested byDendroctonus pseudotsugae.The sex of the adults was determined whenever possible, using the sex characters described by Struble and Carpelan (26). The submental pit present only on the males ofT. virescenswas easy to observe with a 10x hand lens and was very dependable. However, the sex character suggested by the same workers for Esphegeuswas not easy to observe even under a microscope. It was found that the large size of the abdomen of certain clerids usually indicated the condition of a gravid female This character, although not too dependable at times, was used for E.sphegeus, E. lecontei,andT. undatulus.No other secondary sex characteristic could be found. The adults of each species were originally placed in pint jars, and later in clear, plastic boxes, measuring 1 by 2.5 by 3.5 inches. A coarse piece of paper wasglued to the bottom of the box to provide traction for the adults. Usually there was only one pair of each species per container, but occasionally more. This increased the probability of obtaining eggs in instances where sex determinations were not definite; however, there was the disadvantage of increasing mortality since the adults, as well as the larvae, are cannibalistic, especially under crowded conditions. A spiral of blotting paper was made by rolling a triangular piece one to two inches wide at the base. One spiral was placed in each container. It served fairly well as an oviposition site when rolled to about one-quarter inch diameter, and most of the eggs were deposited between the overlapping layers. At times, especially with T virescens,the eggs were laid around the edges of the paper bottom of the box, but these eggs were usually damaged by the activity of the adults. The spiral pieces of paper were examined at various intervals by unrolling the spiral. Portions of the spiral containing eggs were cut to minimize the amount of damage to the eggs. They were then placed in another small plastic box and left there during the incubation period. Living adults of the Douglas-fir beetle were supplied in various numbers and at different intervals of time as food for the ovipositing predators. As soon as eclosion took place, the larvae were removed by means of a camel's-hair brush and placed individually in a 3- by- by ft-inch clear plastic box. Only one larva could be placed in a container because all species are cannibalistic. A piece of blotting paper was placed over the larva to simulate bark, as it was found that the small larvae had difficulty in obtaining a purchase on their prey when this paper was absent. Each larva was fed enough Douglas-fir beetle larvae to insure survival. As in the case of the adults, no attempt was made to record the number of hosts consumed or to maintain a constant feeding schedule. All rearings were made in the laboratory without attempting to control or measure temperature and humidity. Measurements were taken at 15x, 30x, and 90x magnifications, depending on the size of the subject, with a dissecting microscope having a calibrated-micrometer eyepiece. Each micrometer unit at the three magnifications was equivalent to 0.096 mm, 0.034 mm, and 0.017 mm, respectively. All measurements were taken while the subject was immersed in alcohol. Microscope slides were prepared for certain structures in order to make more detailed observations and drawings. Drawings were made by using the same microscope, but with an eyepiece having a 10 by 10 grid. Tracing paper was placed over different scales of graph paper, and the image of the subject trans- ferred from the microscope to the tracing paper by eye with the aid of the grid lines. All drawings were made from preserved specimens by L. N. Kline with the exception of the two ofCoeloides brunneri, which were redrawn from Ryan (21). Terminology in the descriptions follows Knull(11),Peterson (18,19), and Torre-Bueno(27).

Known and Possible Predators and Parasites Past studies concerning the Douglas-fir beetle have reported many species of insects that were supposedly predators or parasites. However, more recent studies have reduced the list of insects believed to cause appreciable mortality. The species which are known predators and parasites of the Douglas-fir beetle and those which are commensals or possibly predaceous or parasitic are listed in this section. The associated arthropods are believed to prey upon the secondary insects associated with the Douglas-fir beetle (Cerambycidae, Bupres- tidae, and Scolytidae), or they may be scavengers. Even some of the species which are listed as being known predators and parasites may cause only a very small percentage of mortality. Known predators and parasites CLASS INSECTA: Coleoptera : Cleridae-Enoclerus sphegeusFab. Enoclerus leconteiWolc Thanasimus undatulusSay Ostomatidae-Temnochila virescens chlorodiaMann. Diptera : Dolichopodidae-MedeteraaldrichiiWh. Medeterasp(nearnigripesLev ) Medeterasp. (nearoregonensisVan Duzee) Lonchaeidae-Lonchaeasp. (nearcorticisTaylor) Lonchaeasn. (nearwatsoniCurran) Hymenoptera: Braconidae-CoeloidesbrunneriVi&. Pteromalidae-RoptroceruseccoptogasteriRatz Cecidostiba burkeiCrawford Cecidostiba dendroctoniAshm Commensals or possible predators and parasites CLASS TNSECTA: Coleoptera :Ostomatidae-Tenebroides sp Histeridae-Undetermined species Staphylinidae-Undetermined species Othniidae-Undetermined species Tenebrionidae-Corticeus sp. Melandryidae-Rushia sp. Colydiidae-Lasconotus sp Hemiptera: Anthocoridae-Lyctocoris sp Diptera: Scenopinidae-Undetermined species Stratiomylidae-Undetermined species Itonididae-Undetermined species Empididae-Undetermined species CLASS ARACHNIDA: ORDER PSEUDOSCORPIONIDA

Field Key to the Insect Larvae of the Known Predators and Parasites The following key is based on mature larvae and is designed for use with a 10x hand lens. No attempt has been made to separate the different instars, eggs, or pupae. Reference can be made to the descriptions of the species if-separation of these stages is needed. Although color is generally undesirable as a primary separation character, it is often the most conspicuous feature of a larva and is therefore useful in field determinations. For this reason, color is used in separating the ostomatids from the clerids.

FIELD KEY

1 Larvae with legs------2 -Larvae without legs------5 2Ventral mouthparts retracted; mesothoracic plates, except for first instar, superficially resembling one plate;all thoracic plates very visible; abdominal segments one to six with dorsal ampullae; apex of mandibles dentate; head, thoracic plates, basal plate and urogomphi very dark brown to black; abdomen white to blue gray at maturity. (Plate XII, Figure A; Plate XIII, Figures B, C, G, and H; Plate XIV, Figure A) ...... ------Temnochila virescens -Ventral mouthparts protracted; mesothoracic plates distinctly separated; all thoracic plates not very visible; abdominal segments without dorsal ampullae ; apex of mandibles entire ;head, thoracic plates, basal plate and urogomphi yellow ocher to dark brown ; abdomen pink to lilac at maturity (Plate III, Figures B, C, and E; Plate VI, Figures J and K; Plate VII, Figure B; Plate IX, Figures D, E, and F)------3 Epicranium with a dorsal tubercle on each side. (PlateI,Figure ti; Plate II, FiguresC andF) ------Enoclerus sphegeus Epicranium without dorsaltubercle. (Plate VI,Figures B and L; Plate VIII, Figure F; Plate IX, FigureC) ------4 Urogomphi somewhat swollen near apex. (PlateVI, FigureH) ------Enoclerus lecontei Urogomphi not swollen nearapex.(Plate IX, FigureA) ------_- ...... Thanasimus undatulus Body slender andcylindrical;with ventralpseudopodia.(PlateXV, Fig- ures B and C) Body crescent-shaped,fusiform,orcyphosomatic;without ventral pseudopodia(Plate XVI,Figures A andD) ------7 6 Body cylindrical, tapering slightly at each end ;internal cylindrical, meta- cephalic and tentorial rodsblack;headsubdivided;a small sclerotized plate on posterior region of head and anterior margin of prothorax; prothoracic spiracles small, circular; caudal pair larger, oval, with two large oval openings(Plate XV,Figure B) ------_------Medetera sp Body wedge-shaped ormuscidiform,tapering gradually to a sharply pointed cephalic end; internal rods dark brown; fused at two locations, branchingcaudally;head notsubdivided;no sclerotized plates on head or prothorax;prothoracic spiracles nine-lobed; caudal pair containing three slits situated at right angles to eachother. (Plate XV, Figure C) ------

------Lonchaea sp. 7Bodyfusiform tocyphosomatic;spiracles located on prothorax and abdominal segments one to eight ;lines between mouthparts and parts of head capsule heavilysclerotized,stripital sclerome between cardo and stripes;seven dorsal protrusile areas in the successive intersegmental areas,the most anterior between the metathoracic and first abdominal segment.(PlateXVI, Figures C and D)....(Braconidae)Coeloides brunneri Body wedge-shaped ormuscidiform,tapering gradually to a sharply end; spiracles located onmesothorax,metathorax and abdominal segments one toseven;lines between mouthparts and parts of head capsule not or feeblysclerotized,cardo and stripes fused or nearlyso,without a conspicuous stipital sclerome between them;without dorsal protrusile areas in the successive intersegmented areas. (PlateXVI, Figures A and B)------Pteromalidae

Description and Illustration of the Egg, Larva, and Pupa of the Known Predators and Parasites Cleridae : Enoclerus sphegeusFabrieius The adult was originally described by Fabricius in 1787 (12, p. 385:16,p. 84), with a more detailed description by Brown in 1957 (20). The mature larva was briefly described and illustrated by Roving (5, pp. 632-633) in 1920. Bedard (1, pp. 33-34) made very sketchy descriptions and illustrations of the egg and mature larva in 1933. In 1957, Reid (20) described the egg and four instars with some drawings. This species is one of the more abundant and widely distributed clerids in the coniferous forests of western North America. It is an important enemy of many species of Scolytidae, principallyDendroc- tonusandIps.Records indicate that the species is found as far south as Mexico and north into British Columbia, with its eastern boundary in Colorado and New Mexico(12, p.150;16,p. 84). Because of its abundance and apparent aggressiveness, this insect can be rated the most important coleopterous predator ofD. pseudotsugae. Briefly, the life cycle with special reference to the Douglas-fir beetle may be summarized as follows : There is only one generation per year, and the emergence of adults is closely synchronized with the emergence of the Douglas-fir beetle in April or May. Mating takes place on and around material being invaded or already infested by the Douglas-fir beetle, and eggs are laid in clusters under scales of the outer bark. The first larval instars apparently enter the cambial region of the tree through the entrance and "ven- tilation" holes constructed by the bark beetles. The larvae develop under the bark, feeding on the eggs, larvae, pupae, and callow adults of its host. All three instars are present throughout the summer. A large percentage of the mature larvae (third instar) emerge in mid- summer and migrate on the outer bark to the base of the tree. Pupal cells are formed in the outer bark or in the duff around the root collar. The clerid overwinters as a prepupal larva. When this species was reared in the laboratory, it appeared not to require diapause. Two generations of this species were reared for identification in the laboratory from adults collected in Idaho and Oregon. Descriptions of immaturestages. The egg, three larval instars, and pupa are described. Egg:(Plate I, Figure A). Average length, 3.1 mm; range in length, 2.5 mm to 3.5 mm. Average width, 0.7 mm; range in width, 0.6 mm to 0.8 mm. Form subcylindrical, with a slightly curved longitudinal axis, tapering slightly at each end, broadest at the middle, a little wider at one end than at the other; margins slightly unequal; chorion smooth, shining, transparent, without ornamentation, becoming slightly rugose and dull as the embryo approaches maturity; color at first pale orange pink, with development gradually becoming bright salmon pink with translucent or opaque white portions at each end and lateral margins. First instar:(Plate I, Figures B to E). Maximum length of larva, 7.30 mm; minimum length, 3.25 mm. Maximum width of body, 1.11 mm; maximum thickness of body, 1.11 mm. Average width of head capsule, 0.69 mm; range of width of head capsule, 0.64 mm

,in E

PLATE I Enoclerus sphegeusFab.: A,egg; B, first instar, lateral view of head; C, dorsal view of head; D, dorsal view of basal plate and urogomphi; E, lateral view of basal plate and urogomphi. to 0.72 mm. Anterior width of prothoracic shield, about 0.72 mm. Width of basal plate of urogomphi, about 0.35 mm. Form orthosomatic; abdomen membranous, pale pink, with a few long, strong, scattered setae; ten abdominal segments with segments four to seven slightly wider; ninth abdominal segment crescentiform, dorsally with a basal plate and paired urogomphi; tenth segment located below the ninth, developed as a locomotive organ, with an ambulatory wart and anal opening; ambulatory wart surrounded anteriorly by four small papillae and posteriorly by one large, liplike lobe; papillae indistinct. Head prognathous, exserted or slightly inserted;dorsal surface somewhat flattened, ventral surface somewhat convex, lateral marginsparallel; as wide as long; with scattered setae, majority same length as mandible; heavily sclerotized, dark brown. Frons triangular, delimited by slightly curved frontal sutures which posteriorly form an acute angle; medially and internally with a well sclerotized, dark brown longitudinal endocarnia, one-half the length of head, branching posteriorly to form a "Y"; two parallel, slightly elevated ridges, one on each side of anterior portion of endocarina. Epicranium dorsally separated by frons into two epicranial halves, ventrally separated by an elongate, rectangular gula; slightly rugose; a very small tubercle located dorsally on each epicronial half near the middle- posterior portion of the frontal sutures. Ocelli located on epicranium behind ventrolateral part of antennal ring;arranged in an anterior row of three and a posterior row of two; rows parallel. Clypeus and labrum lightly sclerotized, yellow ocher, not distinct; anterior margin of labrum with a row of very small setae and posteriorly with a row of four to six longer setae. Antenna lightly sclerotized, yellow ocher; projecting from an antennal ring; extending beyond anterior margin of labrum; basal membrane large, whitish, transparent, and enclosing about one-half of basal segment; basal segment twice as long as second segment; second segment with a small appendix, three setae around distal margin; apical segment cylindrical, about three-fourths the length of basal segment, apex with one long seta surrounded by three very short setae. Mandible subtriangular, apex pointed, about three-fourths the length of frons, width at base slightly more than one-half length of mandible; retinaculum slightly closer to apex than to base of mandible; two short setae on the lateral mandibular face; with a longitudinal groove on the ventral surface. Ventral mouthparts protracted with distal half directed obliquely upwards; lightly sclerotized, yellow ocher. Prothorax dorsally with a tergal shield or plate; heavily sclerotized, dark brown; anterior margin straight, curving ventrocephalad; rounded, posteriorly oblique side margins which end at the dorsal notch; a longitudinal endocarnia starting from dorsal notch and continuing medially and anteriorly for three-fourths of the length of prothoracic plate, dark brown; long setae around outer margin with shorter setae internally. Ventrally, with a pair of subtriangular, presternal plates, one on each side of a narrow, lanceolate, sternal plate;all plates lightly sclerotized and very light yellow ocher or light brown. Mesothorax dorsally with two, subtriangular plates; one small seta on each; ventrally with a very small, obtuse plate posterior to prothoracic lanceolate plate; a small, oblong plate posterior to obtuse plate. Metathorax dorsally with two pentagonal plates ; plates smaller and farther apart than mesothoracic plates; one small seta on each; ventrally with a small, oblong plate. Mesothoracic and metathoracic dorsal plates lightly sclerotized and light brown, less distinct on the living specimen than the prothoracic dorsal plate. Thoracic legs five-segmented, no free claws. Spiracles annular-biforous, located laterally on the mesothorax and abdominal segments one to eight; mesothoracic spiracle slightly larger than abdominal spiracles; metathoracic spiracle, rudimentary. Basal plate of urogomphi lying at an angle on ninth abdominal segment; not well defined, fading into and surrounded by a lightly sclerotized, yellow ocher, crescent-shaped sclerite; length about one- half as long as frons, slightly wider than long; lightly sclerotized, dark brown ocher. Urogomphi subconical; directed slightly upwards, markedly divergent at apex, apex not recurved;outer margins unequal, inner margins equal; about as long as length of basal plate; heavily sclerotized, dark brown. A few scattered setae on basal plate, about as long as width of basal plate;setae on urogomphi more numerous and up to three times as long as urogomphi. Second instar:(Plate II, Figures A to C). Maximum length of larva, about 17 mm; minimum length, about 14 mm. Maximum width of body, about 1.5 mm; maximum thickness of body, about 1.3 mm. Average width of head capsule, 0.99 mm; range of width of head capsule, 0.80 mm to 1.02 mm. Anterior width of prothoracic shield, about 1.04 mm. Width of basal plate of urogomphi, about 0.69 mm. Abdomen bright pink to light lilac color at maturity; dorsally the lilac color appearing as a mottled pattern with intervenientareas and ventral side being whitish lilac;setae more abundant. Head dark brown to black; dorsal epicranial tubercles becoming more prominent; apical segment of antenna about same length as second segment; anterior row of ocelli subparallel to slightly curved in relation to posterior row. Prothoracic shield darker brown. Basal plate of uro-

13 A 0.0 MM B

0.5 MM

PLATE 11 Enoclerus sphegeusFab.: A, second instar, dorsal view of basal plate and urogomphi ;B, second instar, lateral view of basal plate and urogomphi ;C, second instar, lateral view of head; D, third instar, dorsal view of basal plate and urogomphi ;E, third instar, lateral view of basal plate and urogomphi ; F, third instar, lateral view of head. gomphi well defined; anterior margin with a thin, subparallel, lightly sclerotized sclerite; length slightly less than one-half as long as frons, wider than long; heavily sclerotized, dark brown ocher. Urogomphi subconical; directed distinctly upwards, slightly divergent at apex, apex sharply recurved and turned slightly inwards; outer margins unequal, inner margins slightly unequal; about two-thirds to same length as basal plate; heavily sclerotized, dark brown. Setae more numerous on basal plate and urogomphi, scattered, about as long as width of basal plate; few small. Third instar:(Plate IT, Figures D to F; Plate III, Figures A to G; Plate IV, Figures A and B; Plate V, Figure D). Maximum length of larva, about 22 mm; minimum length, about 15 mm. Maxi- mum width of body, about 3.0 mm; maximum thickness of body, about 2.7 mm. Average width of head capsule, 1.53 mm; range of width of head capsule, 1.40 mm to 1.72 mm. Anterior width of prothoracic shield, about 1.77 mm. Width of basal plate of urogomphi, about 1.19 mm. Abdomen dorsally light lilac to deep lilac color at pupation;ventrally more bluish gray;lilac color appearing as a mottled pattern with intervening areas very light blue; thorax more pinkish;setae very numerous and of varying lengths. Head dark brown to black. Frons with two circular, cushionlike elevations, one at base of each mandible; also with four parallel, slightly elevated ridges, two on each side of endocarina; posterior portion of frons rugose. Dorsal epicranial tubercles very large and prominent; epicranial halves very rugose; anterior and posterior rows of ocelli subparallel. Labrum and clypeus distinct. Antenna brown; basal segment about two to three times as long as second segment; apical segment about two-thirds the length of second segment. Metathoracic plates each bearing one long centrally located seta surrounded by four smaller setae. Basal plate of urogomphi about one-half the length of frons, twice as wide as long. Urogomphi not divergent at apex, apex turned more inwards; outer and inner margins more nearly equal; about the length of basal plate; heavily sclerotized, dark brown with apex and margins black; setae numerous, a few small, with remainder as long as urogomphi. The four small, anterior papillae of ambulatory wart very distinct. Pupa:(Plate V, Figure B). Total length 8 mm to 10 mm; typically exarate; abdomen with nine visible segments dorsally and seven ventrally; head, thorax, and abdomen pink with white appendages, abdomen becoming lilac at maturity; a few, small, strong, scattered setae on head, legs, and abdomen; antennae bent downward along pleura; lateral spiracles on abdominal segments one to seven; apices of wings subequal, extending to middle of fourth abdominal segment;

15 q

05 MM

PLATE III Enoclerus sphegeus Fab., third instar: A, dorsal view of head; B, dorsal view of mandible;C, ventral view of mandible;D, mesothoracic leg ; E, ventral view

of head; F, dorsal view ofantenna;G, abdominal spiraclelp, T r - },x .\'A

PLATE IV

Enoclerus sphegeus Fab., thirdinstar:A, dorsal view of complete larva; ventral view of complete larva PLATE V A, ventral view of pupa ofEnoclerus leconteiWolc.; B, ventral view of pupa ofEnoclerus sphegeusFab. ;C, third instar, lateral view of complete larva of Enoclerus sphegeusFab. ;D, ventral view of pupa ofThanasimus undatulus Say.

18 apices of prothoracic tarsi extending to apex of thorax, those of mesothoracic tarsi to apex of second abdominal segment, those of metathoracic tarsi to apex of fifth abdominal segment; anal cerci present. Cleridae:Enoclertiis leconteiWolcott The adult was originally described by LeConte in 1861 asClerus nigriventrisfrom specimens collected near Fort Colville, Washington, and from the Bitterroot Valley, Montana (2, p. 33;17).Schenkling in 1910 placed LeConte'snigriventrisin the genusThanasimusand called itT. nigriventrisThe species was redescribed by Wolcott as Clerus leconteiin 1910.1 Wolcott (4) stated, "The namenigriventris is preoccupied inClerus by C. nigriventrisBlanchard (an Argentina species described in 1842). Should the species be reassigned to the genusClerus,the specific name ofleconteiis suggested for it." In the same year, the genusCleruswas changed toEnoclerus (12). Boving (4) in 1928 called the cleridEnoclerus leconteiWolc. The clerid was referred to asThanasimus lecontei(Wolc.) by Person in 1940(17),but he recently concluded that this was a mistaken identification.2 Blackwelder in 1945 cited the species asT. nigriventris (Lee.) and gaveleconteiWolc. as a synonym.' Apparently, Black- welder missed Blanchard's speciesnigriventrisbecause there is no mention ofitin his checklist. The species was calledEnoclerus leconteiWolc. by Wolcott in his catalog of 1947(28, p.79). Black- welder (3, p. 14) in 1948 followed Wolcott. Apparently the correct name at present isEnoclerus leconteiWolcott. The mature larva was described and illustrated by Boving (4) in 1928 from material collected fromIpsgalleries near Coeur d'Alene, Idaho. The following is the first known description and illustration of all immature stages of this species. One generation was reared in the laboratory from adults collected in Oregon. The known distribution of this species extends from British Columbia to Michigan and south to Guatemala(16,p. 84). It has been reported mainly as an enemy of bark beetles infestingPinus. Bedard (1, p. 36) reports that it was associated with the Douglas-fir beetle,Dendroctonus pseudotsugae,in Montana. He states, "The scarcity of this insect in Douglas-fir prohibits it from being of any great value in the control of the Douglas-fir beetle." The clerid was not observed to be associated with the Douglas-fir beetle in southern

1 Personal communication, P. 1. Spangler, United States National Museum, Washington, D C 2 Personal communication, H. L Person, Foreign Forestry Services, Wash- ington, D. C. Idaho. As far as is known, there has been no earlier report of the presence ofthisspecies in association withD. pseudotsugaein western Oregon. Again, it is believed that E.leconteiis of no great importance in controlling the Douglas-fir beetle in this region. The clerid seems to be more abundant in association with the smaller bark beetles, such asScolytus unispinosusLec. and various species of the genusPseudohylesinusIts life cycle appears to be similar to E. sphegeusin Oregon. Descriptions of immaturestages. The egg, three larval instars, and pupa are described. Egg:(Plate VI, Figure A). Average length, 1.3 mm; range in length, 1.2 mm to 1.6 mm. Average width, 0.4 mm; range in width 0.3 mm to 0.4 mm. Form subcylindrical, with a slightly curved longitudinal axis, tapering slightly at each end, broadest at the middle, a little wider at one end than at the other; margins equal; chorion smooth,shining,transparent,withoutornamentation,becoming slightly rugose and dull as the embryo approaches maturity; color at first pale pink, with development gradually becoming brighter pink with translucent or opaque white portions at each end and lateral margins. First instar:(Plate VI, Figures B to E). Maximum length of larva, 2.18 mm; minimum length, 1.34 mm. Maximum width of body, 0.37 mm; maximum thickness of body, 0.34 mm. Average width of head capsule, 0.27 mm; range of width of head capsule, 0.23 mm to 0.30 mm. Anterior width of prothoracic shield, about 0.29 mm. Width of basal plate of urogomphi, about 0.18 mm. Form orthosomatic; abdomen membranous, very pale pink, with long, strong, scattered setae;ten abdominal segments with segments four to six slightly wider; ninth abdominal segment somewhat crescentiform, dorsally with a basal plate and paired urogomphi; tenth segment located below the ninth, developed as a locomotive organ, with an ambulatory wart and anal opening; papillae of ambulatory wart indistinct. Head prognathous, exserted or slightly inserted; dorsal surface flattened to slightly concave, ventral surface and lateral margins slightly convex; as wide as long; with scattered setae, majority same length as mandible; heavily sclerotized, light brown. Frons triangular, delimited by slightly curved frontal sutures which posteriorly form an acute angle; medially and internally with a well sclerotized, dark brown longitudinal endocarnia, two-thirds length of head, branching posteriorly to form a "Y." Epicranium dorsally separated by frons into two epicranial halves, ventrally separated by an elongate, rectangular gula; epicranial halves without tubercles. Ocelli located on Aco1MM

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PLATE VI Enocicrus lecontciWole.: A,egg; B, first instar, lateral view ofhead;C, first instar,dorsal view of head; D, first instar, lateral view of basal plate and urogomphi; E, first instar, dorsal view of basal plate and urogomphi ; F, second instar, lateral view of basal plate andurogomphi;G, secondinstar,dorsal view of basal plate andurogomphi;H, thirdinstar,dorsal view of basal plate and urogomphi;I, thirdinstar,lateral view of basal plate andurogomphi;J, third instar, ventral view ofmandible;K, thirdinstar,dorsal view of mandible; L, thirdinstar,lateral view of head.

21 epicranium behind ventrolateral part of antennal ring; arranged in an anterior row of three and a posterior row of two; rows parallel. Clypeus and labrum not distinct; anterior margin of labrum with a row of small setae. Antenna slightly sclerotized, light yellow ocher ; projecting from an antenna] ring; extending beyond anterior margin of labrum;basal membrane large, whitish, transparent, and enclosing about one-half of basal segment; basal segment about three to four times as long as second segment; second segment with a small appendix, three setae around distal margin; apical segment cylindrical, same length as second segment, apex with one long seta. Mandible subtriangular, apex pointed, about three-fourths the length of frons, as wide at base as long;retinaculum near middle of mandible; two short setae on the lateral mandibular face; with a longitudinal groove on the ventral surface. Ventral mouthparts protracted with distal half directed somewhat obliquely upwards; lightly sclerotized, yellow ocher. Prothorax dorsally with a tergal shield or plate; heavily sclerotized, light brown; anterior margin straight, curving ventrocephalad; rounded, posteriorly oblique side margins which end at the dorsal notch;a longitudinal endocarnia starting from dorsal notch and continuing medially and anteriorly three-fourths length of prothoracic plate, (lark brown; with scattered setae. Ventrally, with a pair of subtriangular, presternal plates, one on each side of a narrow, lanceolate, sternal plate;allplates lightly sclerotized and very light yellow ocher; not too distinct on living specimen. Mesothorax dorsally with two subtriangular plates; one small seta on each; ventrally with a very small, obtuse plate posterior to prothoracic lanceolate plate; a small, oblong plate posterior to the obtuse plate; ventral plates not distinct on living specimen. Metathorax dorsally with two subquadrangular plates;plates smaller and farther apart than mesothoracic plates; one small seta on each; ventrally with a small, nondistinct, oblong plate. Meso- thoracic and metathoracic dorsal plates lightly sclerotized and light brown, less distinct on the living specimen than the prothoracic dorsal plate. Thoracic legs five-segmented, no free claws. Spiracles annular-biforous, located laterally on the mesothorax and abdominal segments one to eight; mesothoracic spiracle slightly larger than abdominal spiracles; metathoracic spiracle, rudimentary. Basal plate of urogomphi lying at an angle on ninth abdominal segment; surrounded by a lightly sclerotized, yellow ocher, crescent- shaped sclerite; length slightly shorter than frons, wider than long; lightly sclerotized, light brown ocher. Urogomphi subconical; strongly divergent at apex, apex not recurved;outer and inner margins unequal; slightly longer than length of basal plate; heavily sclerotized, light brown. A few scattered setae on basal plate, about as long as basal plate; setae on urogomphi more numerous and up to four times as long as urogomphi. Second instar:(Plate VI, Figure F and G). Maximum length of larva, about 5 mm; minimum length, about 4 mm. Maximum width of body, about 0.80 mm; maximum thickness of body, about 0.77mm. Average width of head capsule, 0.47 mm; range of width of head capsule, 0.37 mm to 0.52 mm. Anterior width of prothoracic shield, about 0.52 mm. Width of basal plate of urogomphi, about 0.40 mm. Abdomen bright pink; setae more abundant. Head and prothoracic shield dark brown. Basal plate of urogomphi well defined; surrounded by a narrow, lightly sclerotized sclerite; length about two- thirds as long as frons, slightly wider than long; somewhat sculptured with fine subparallel depressions running length of plate; heavily sclerotized, light brown ocher. Urogomphi subconical;swollen at base, not divergent at apex; apex not recurved but turned slightly inwards; outer margins unequal, inner margins equal; length about one-half as long as basal plate; heavily sclerotized, dark brown. Setae scattered, slightly longer than length of basal plate. Third instar:(Plate VI, Figures H to L; Plate VII, Figures A to E). Maximum length of larva, about 11 mm; minimum length, about 8 mm. Maximum width of body, about 1.6 mm; maximum thickness of body, about 1.3 mm. Average width of head capsule, 0.80 mm; range of width of head capsule,0.77mm to 0.82 mm. Anterior width of prothoracic shield, about 0.91 mm. Width of basal plate of urogomphi, about 0.75 mm. Abdomen bright pink, becoming lilac color starting from the posterior end, very dark lilac at pupation; dorsally very dark mottled lilac pattern, ventrally more gray lilac; thorax more reddish brown; setae very numerous and about the same length. Head very dark brown to black. Frons with two circular, cushion-like elevations, one at base of each mandible; also with two parallel, slightly elevated ridges, one on each side of endocarnia. Anterior and posterior rows of ocelli subparallel. Labrum and clypeus distinct. Basal plate of urogomphi surrounded by a narrow, lightly sclerotized sclerite;slightly less than one-half the length of frons, twice as wide as long; somewhat sculptured with irregular shaped depressions scattered over plate; dark brown. Urogomphi subconical to subtriangular;swollen from base to near apex; apex turned sharply inwards; outer and inner margins unequal; length about two-thirds as long as basal plate; very dark brown to black. Setae scattered, a few very small with remainder about one-half to three-

23 PLATE VII Enoclerus leconteiWolc, thirdinstar: A, dorsal view of head ; B, ventral view of head ;C, dorsal view of complete larva ; D, ventral view of complete larva lateral view of complete larva fourths width of basal plate. Ambulatory wart surrounded anteriorly by four small papillae and posteriorly by one large lobe and anal opening; papillae and lobe very distinct. Pupa:(Plate V, Figure A). Total length, about 6 mm; typically exarate; abdomen with nine visible segments dorsally and seven ventrally; head and thorax pink, abdomen dark pink with mottled lilac areas; appendages white to gray; a few small, strong, scattered setae on head, legs, and abdomen; antennae bent downward along pleura; apices of wings equal, extending to apex of fourth abdominal segment; apices of prothoracic tarsi extending to apex of thorax, those of mesothoracic tarsi to middle of third abdominal segment, those of metathoracic tarsi to apex of fifth abdominal segment; anal cerci present. Cleridae :Thanasimus undatulusSay The adult of this species was described by Say in 1835 (12, p. 149) asThanasimus undatulus.Leng in 1920(12, p.149) listed the species asT. undulatusSay. In 1927(13,p. 28) it was reported that undulatuswas a typographical error forundatulus.This insect was mistakenly identified asT. dubius(Fab.) by Bedard(1,p. 28). The forms of the species appear different throughout its geographical distribution. Barr states, "I prefer for the present to call everything T. undatulus."3 It has been reported that the insect ranges from Alaska to New Mexico and eastward to New Hampshire(12, p.149;16,p. 82). The clerid occurs occasionally in association with the Douglas-fir beetle in Montana, Idaho, and Oregon; its preferred prey are smaller bark beetles such as various species ofPseudohylesinus, Scolytus,etc. The life cycle of this species appears to be similar to that of E.sphegeus, with the exception that the larvae do not migrate to the base of the tree. Two generations were reared in the laboratory from adults collected in Idaho. The following is the first known description and illustration of all of the immature stages. Descriptions of immaturestages. The egg, three larval instars, and pupa are described. Eggs:(Plate VIII, Figure E). Average length, 2.3 mm; range in length, 2.0 mm to 2.2 mm. Average width, 0.5 mm; range in width, 0.5 mm to 0.6 mm. Form subcylindrical, with a slightly curved longitudinal axis, tapering slightly at each end, broadest at the middle, one end slightly wider than the other, margins equal; chorion smooth, 'Personal communication, W. F. Barr, University of Idaho, Moscow. A 0.25 mm C , 0.5 YY

IYY

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PLATE VIII Thanasimus undatulus Say :A, firstinstar, dorsal view of basal plate and urogomphi ; B, first instar, lateral view of basal plate and urogomphi ;C, second instar, dorsal view of basal plate and urogomphi ; D, second instar, lateral view of basal plate and urogomphi; E, egg; F, first instar, lateral view of head; G, first instar, dorsal view of head; H, third instar, dorsal view of head. shining,transparent, without ornamentation;color opaque white throughout development. First instar:(Plate VIII, Figure A, B, F, and G). Maximum length of larva, 3.77 mm; minimum length, 2.63 mm. Maximum width of body, 0.53 mm; maximum thickness of body, 0.50 mm. Average width of head capsule, 0.36 mm; range of width of head capsule, 0.35 mm to 0.38 mm. Anterior width of prothoracic shield, about 0.40 mm. Width of basal plate of urogomphi, about 0.27 mm. Form orthosomatic; abdomen membranous, very pale pink, with a few, long, strong, scattered setae; ten abdominal segments with segments four to seven slightly wider; ninth abdominal segment somewhat crescentiform, dorsally with a basal plate and paired urogomphi; tenth segment located below the ninth, developed as a locomotive organ, with an ambulatory wart and anal opening; ambulatory wart surrounded anteriorly by four small papillae and posteriorly by one large, liplike lobe; papillae indistinct. Head prognathous, exserted or slightly inserted; dorsal surface flattened, ventral surface and lateral margins slightly convex; as wide as long; with scattered setae, majority same length as mandible; heavilysclerotized,dark brown. Frons triangular,delimited by slightly curved frontal sutures which posteriorly form an acute angle; medially and internally with a well sclerotized, dark brown, longitudinal endocarnia, one-third length of head, branching posteriorly to form a "Y." Epicranium dorsally separated by frons into two epicranial halves, ventrally separated by an elongate, rectangular gula; epicranial halves without tubercles. Ocelli located on epicranium behind ventrolateral part of antennal ring; arranged in an anterior row of three and a posterior row of two; rows subparallel. Clypeus and labrum lightly sclerotized, yellow ocher, not distinct; anterior margin of labrum with a row of very small setae. Antenna lightly sclerotized, yellow ocher; projecting from an antennal ring; extending beyond anterior margin of labrum; basal membrane large, whitish, transparent, and enclosing about one-fourth of basal segment; basal segment about three times as long as second segment; second segment with a small appendix, three setae around distal margin; apical segment cylindrical, about twice the length of second segment, apex with one long seta. Mandible subtriangular, apex pointed, about three- fourths the length of frons, width at base about two-thirds length of mandible; retinaculum slightly closer to apex than to base of mandible; two short setae on the lateral mandibular face; with a longitudinal groove on the ventral surface. Ventral mouthparts protracted with distal half directed obliquely upwards; lightly sclerotized, yellow ocher.

27 Prothorax dorsally with a tergal shield or plate; heavily sclerotized, light brown, anterior margin straight, curving slightly ventrocaudad; rounded, posteriorly oblique side margins which end at the dorsal notch; a longitudinal endocarnia starting from dorsal notch and continuing medially and anteriorly three-fourths length of prothoracic plate, dark brown; with scattered setae. Ventrally, with a pair of triangular, presternal plates, one on each side of a narrow, lanceolate, sternal plate;all plates lightly sclerotized and very light yellow ocher. Mesothorax dorsally with two subtriangular plates; three small seta on each; ventrally with a very small, obtuse plate posterior to prothoracic lanceolate plate; a small, oblong plate posterior to obtuse plate. Metathorax dorsally with two subquadrangular plates;plates smaller and farther apart than mesothoracic plates; two small setae on each; ventrally with a small, nondistinct, oblong plate. Mesothoracic and metathoracic dorsal plates lightly sclerotized and light brown, less distinct on the living specimen than the prothoracic dorsal plate. Thoracic legs five-segmented, no free claws. Spiracles annular-biforous, located laterally on the mesothorax and abdominal segments one to eight; mesothoracic spiracle slightly larger than abdominal spiracles; metathoracic spiracle rudimentary. Basal plate of urogomphi lying horizontally on ninth abdominal segment; anterior margin at times not well defined; two longitudinal grooves extending length of plate; length about two-thirds as long as frons, wider than long;lightly sclerotized, light brown ocher. L rogomphi subcylindrical; markedly divergent at apex, apex not recurved; outer margins slightly unequal, inner margins equal; about two-thirds length of basal plate;heavily sclerotized, dark yellow brown. A few scattered setae on basal plate, about as long as width of basal plate; setae on urogomphi more numerous and up to three times as long as urogomphi. Second instar:(Plate VIII, Figures C and D). Maximum length of larva, about 13 mm; minimum length, about 8 mm. Maximum width of body, about 2.0 mm; maximum thickness of body, about 1.8 mm. Average width of head capsule, 0.63 mm; range of width of head capsule, 0.60 mm to 0.68 mm. Anterior width of prothoracic shield, about 0.69 mm. Width of basal plate of urogomphi, about 0.56 mm. Abdomen bright pink to light lilac at maturity; lilac color starting to appear at posterior end and progressing anteriorly; dorsally the lilac color appearing as a mottled pattern with intervenient areas and ventral side light blue;setae more abundant. Head and prothoracic shield dark brown. Basal plate of urogomphi lying at

28 an angle on ninth abdominalsegment;welldefined;surrounded by a lightlysclerotized,light brownsclerite;a largedepression,centrally located and extending length'ofplate;length about as long as frons, wider than long; heavilysclerotized,dark brown ocher. Urogomphi subcylindrical;directed distinctly upwards, not divergent at apex, apex slightlyrecurved;outer marginsunequal,inner margins equal; slightly more than one-half the length of basal plate; heavily sclerotized, darkbrown.Setae more numerous on basal plate and urogomphi,scattered,about as long as width of basal plate; few small. Third instar: (PlateVIII,FigureH;Plate IX, Figures A to F;Plate X, FiguresA to Q.Maximum length of larva, about 19 mm; minimumlength,about 11 mm. Maximum width of body, about 2.8 mm; maximum thickness ofbody,about 2.7 mm. Average width of head capsule, 1.12 mm; range of width of head capsule, 1.09 mm to 1.19 mm. Anterior width of prothoracic shield, about 1.24 mm. Width of basal plate ofurogomphi,about 0.99 mm. Abdomen dorsally lilac to deep lilac color atpupation;ventrally more bluish;lilac color appearing as a mottled pattern with intervenient areas light blue; thorax morewhitish;setae very numerous and of varying lengths. Head dark reddish brown to black. Frons with two circular,cushionlikeelevations,one at base of each mandible ;also with fourparallel,slightly elevatedridges,two on each side of endocarnia.Posterior portion of head slightlyrugose.Labrum and clypeusdistinct.Antennabrown;basal membrane enclosing about one-half to two-thirds of basalsegment;apical segment about as long as second segment. Basal plate of urogomphi with several depressions extending length ofplate;length three-fourths the width ofplate,and slightly shorter than frons; dark reddish brown. Uro- gomphisubcylindrical;apex sharply recurved and turned inwards; outer and inner marginsunequal;about one-half the length of basal plate;very dark brown toblack;setaenumerous,few small, with remainder slightly less than length ofurogomphi.Four small anterior papillae of ambulatory wart very distinct. Pupa: (PlateV,FigureD).Totallength,about 10mm;typically exarate; abdomen with nine visible segments dorsally and seven ventrally;head and thorax pinkish, appendages white, abdomen dark lilac; a few small,strong,scattered setae on head,legs,and abdomen; antennae bent downward alongpleura;lateral spiracles on abdominal segments one toseven;apices of wingssubequal,extending between the fourth and fifth abdominalsegments;apices of prothoracic tarsi extending to apex ofthorax,those of mesothoracic tarsi to apex of third abdominalsegment,those of metathoracic tarsi to apex of fifth abdominalsegment;anal cerci present. A

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PLATE IX Thanasimus undatulus Say, third instar :A, dorsal view of basal plate and urogomphi; B, lateral view of basal plate and urogomphi; C, lateral view of head; D, ventral view of head; E, dorsal view of mandible; F, ventral view of mandible. I

IYM PLATE X C, lateral view of complete larva Say, third instar: A, dorsal view of complete larva; B, Thanasimus undatulus ventral view of complete larva ; a

Ostomatidae :Temnochila virescens chlorodia Mannerheim Temrwochila zirdsce#&s was' described by Fabricius in 1775 (12,, p. 193). A subspecies, T. virescens chlorodia, was describedin 1$431 by Mannerheim (12, p. 193). The mature larva of T. vi.rescens Fab. Was illustrated by Roving and Craighead in 1930 (6, pp.272-274). Bedard (1, pp. 38-39) and Struble (23, pp. 100-101) made some very. general descriptions of the mature larva. Records indicate that T. virescens is. found in the eastern states, while T. viresceis ch rod'i is found in the western states (12, p. 193). This species is very abundant and widely distributed in coniferous forests of North America. It is an important enemy of many species of Scolytidae, principallyDendroctonusandIps.Because of its abundance and apparent aggressiveness, this insect can be rated very close toEnoclerus sphegusin importance as a predator of D. pseudotsugaein the Intermountain Region. Its importance as a predator ofD. pseudotsugaein the Pacific Northwest is less. Here the species is less abundant and seems to be somewhat aggregated in its distribution through the forest. It has been observed that the insect prefers more exposed, warmer sites of windthrown Douglas- fir in western Oregon. The life cycle ofT. virescensfollows closely that of the clerids with the exception that the larvae do not migrate. The following is the first known description and illustration of the immature stages of this species. The insect was found to be more difficult toI rear in the laboratory than the clerids. Apparently, the s quantity of food, temperature, and humidity are very important in the development of the larvae. The number of times the larvae molted varied considerably, but the maximum number of instars observed was seven. There was a gradual increase in size of most structures, especially the head capsule, until the seventh molt. At that time the larva were about the size of the fifth instars, and this fact may indicate that the usual number of instars under field conditions is around five or six. There were no major morphological changes after the third instar. Therefore, the drawings of the sixth instar represent the third to sixth instars with the exception of change in size. No pupae were obtained. Two generations were reared in the laboratory from adults collected in Idaho and Oregon. Descriptions of immature stages. The egg and six larval instars are described. Egg:(Plate XI, Figure A). Average length, 2.3 mm; range in length, 2.2 mm to 2.5 mm. Average width, 0.4 mm; range in width, 0.4 mm to 0.5 mm. Form cylindrical, with a slightly curved longi-

I t

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01;

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PLATE XI Temnochila virescens chlorodia Mann :A, egg ;B, first instar, lateral view of head; C, first instar, dorsal view of head and thorax; D, first instar, dorsal view of basal plate and urogomphi ;E, first instar, lateral view of basal plate and urogomphi ;F, second instar, lateral view of basal plate and urogomphi

33 tudinal axis, ends rounded; chorion smooth, dull, transparent, without ornamentation; color very pale, orange pink throughoutdevelop- ment. First instar: (Plate XI, Figures B to E). Maximum length of larva, 3.42nun;minimumlength,2.45mm.Maximum width of body,0.50 mm; maximum thickness of body, 0.44 mm. Aver- age width of head capsule. 0.42 mm; range ofwidth of head capsule.0.40 nun to0.43 mm. Anterior width ofprothoracic plates,about 0.46nom. Widthofbasalplateofurogomphi, about 0.28 mm. Form orthosomatic; abdomen membranous, gray pink;setae strong and scattered, more abundant on ventralside than dorsal side, consisting of three lengths; all setae surrounded by a very small, black papilla, the longer setae raised on asmall chalaza; ten abdominal segments with segments two to seven slightlywider; segments without dorsal ambulatory warts; ninth abdominal segment somewhat crescentiform, dorsally with a basal plate and paired urogomphi; tenth segment located below the ninth, developed as a locomotive organ, with an ambulatory wart and anal opening; ambulatory wart surrounded anteriorly by six small papillae and posteriorly by one large, liplike lobe; papillae indistinct. Head prognathous, exserted or slightly inserted: dorsal surface with small, scattered grooves, ventral surface somewhat flattened. lateral margins parallel; about as wide as long: scattered setae small except for a few along the lateral margins of the epicraniumwhich are about as long as the mandible; heavily sclerotized, (larkreddish brown. Frons triangular, delimited by slightly curved frontal sutures which posteriorly form an acute angle: medially and internally with a well sclerotized, (lark brown, longitudinalendocarnia, about one- third length of head, branching posteriorly to form a "Y." Epi- cranium dorsally separated by frons into two epicranialhalves, ventrally separated by the retracted mouthparts and it short, broad },rata surrounded by a pair of paragula: rugose; epicranial halves without tubercles. Ocelli located on epicranium behind ventrolateral part of antennal ring; arranged in an anterior row of three and a posterior row of two, rows subparallel;a single ocellus located ventrad of the two rows ofocelli.Clypeus and labrum lightly sclerotized, yellow ocher, notdistinct;anterior margin of labrum with a row of very small setae. Antenna lightly sclerotized, yellow Ocher; projecting from an antennal ring; extending beyond anterior margin of labrum: basal membrane large, whitish, transparent, and enclosing about one-fourth of basal segment; basal segment one-half the length of second segment: second segment with a small appendix. three setae around distal margin; apical segment cyclindrical, about

34 three-fourths the length of second segment, apex with one long seta surrounded by two very short setae. Mandible subtriangular, apex dentate with distal ends pointed, about two-thirds the length of frons, width at base about two-thirds the length of mandible; retinaculum situated near the middle of mesal edge of mandible; a penicillus on the edge of a pseudomolar; mesal edge of mandible coarsely serrate; ventral surface somewhat grooved along the distal-mesal edge; two short setae on the lateral mandibular face. Ventral mouthparts retracted with distal half either horizontal or directed slightly down- wards; lightly sclerotized, yellow ocher. Prothorax dorsally with two tergal shields or plates, superficially resembling one plate; heavily sclerotized, anterior-mesal portion of plates light brown with the lateral margins and posterior portions dark brown to black;anterior margins straight, curving ventrocephalad; rounded, posteriorly oblique side margins which end at the dorsal notch; a shallow depression running the length of each plate; long setae around outer margins, with shorter setae internally; a pair of very small, black, subtriangular plates posterior to the large prothoracic plates. Ventrally, three pairs of subtriangular, presternal plates, one pair on each side of a narrow, lancelolate, sternal plate; all plates lightly sclerotized, light brown. Mesothorax dorsally with two, distinctly separated subquadrangular plates; heavily sclerotized, dark brown to black with anterior margin light brown; seven small setae on each plate; a pair of very small, black, subtriangular plates posterior to large, mesothoracic plates. Ventrally, spatulate plate posterior to the prothoracic lanceolate plate, lightly sclerotized, light brown; an oblong plate posterior to the spatulate plate, lightly sclerotized, yellow ocher, not too distinct on living specimen. Metathoraxdorsallywith twosubtriangularplates;plates smaller and farther apart than mesothoracic plates; four small setae on each; heavily sclerotized, dark brown to black. Ventrally with a small, obtuse plate; an oblong plate posterior to obtuse plate; lightly sclerotized, yellow ocher, not too distinct on living specimen. Thoracic legs five-segmented, no free claws. Spiracles annular-biforous, located laterally on the mesothorax and abdominal segments one to eight; mesothoracic spiracle slightly larger than abdominal spiracles; metathoracic spiracle, rudimentary. Basal plate of urogomphi lying at an angle on ninth abdominal segment; well-defined; anterior margin sinuous; plate with two light brown tubercles, each with one seta; length about one-halfas long as frons, twice as wide as long; heavily sclerotized, dark brown to black. Urogomphi subcylindrical; directed slightly upwards, slightly

35 divergent at apex; apex not recurved, circular in cross-sectional view; outer and inner margins equal; about one-half the length of basal plate; heavily sclerotized, brown. Scattered setae on basal plate, a few small with remainder slightly longer than abdominal setae; setae on urogomphi scattered, few small, with remainder up to eight times as long as urogomphi. Second instar:(Plate XI, Figure F; Plate XII, Figures A and B). Maximum length of larva, about 5.40 mm; minimum length, about 3.69 mm. Maximum width of body, about 0.77 mm; maximum thickness of body, about 0.64 mm. Average width of head capsule, 0.72 mm; range of width of head capsule, 0.60 mm to 0.89 mm. Anterior width of prothoracic shields, about 0.80 mm. Width of basal plate of urogomphi, about 0.57 mm. Anterior margin of prothoracic plates straight, curving ventrocaudad. Mesothorax dorsally with two subquadrangular plates;plates not very close together, superficially resembling one plate; anterior margin of plates with a lightly sclerotized, yellow ocher sclerite; each plate with five small setae. Basal plateof urogomphi with the anterior-mesal portion of margin sinuous; lateral margins rounded, curving ventrocaudad; length about three-fourths as long as frons, width nearly twice the length; setae more numerous than on urogomphi; dark brown to black. Urogomphi subconical; not divergent at apex; apex turned sharply upwards and slightly inwards; outer margin unequal, inner margin equal; one- half the length of basal plate; with small setae, a few up to seven times as long as urogomphi; brown with apex black. Third instar:Average length of larva, about 10 mm. Maximum width of body, about 2.01 mm; maximum thickness of body, about 1.68 mm. Average width of head capsule, 1.30 mm; range of width of head capsule, 1.22 mm to 1.37 mm. Anterior width of prothoracic shields, about 1.53 mm. Width of basal plate of urogomphi, about 1.10 mm. Head capsule ovate; posterior portion of endocarnia of head branching to form a small rectangle;posterior portion of epicranial halves nearly meeting each other. Basal segment of antenna about three-fourths the length of second segment; apical segment very short. Clypeus and labrum light brown, distinct. Abdomen becoming blue white. Dorsal ambulatory warts present on abdominal segments two to seven;papillae on second abdominal, ambulatory wart not too distinct or numerous; papillae on remaining ambulatory warts more distinct and numerous, arranged in two rows curving ventrocephalad; few papillae scattered on lateral margins. Ventral prothoracic plates light brown, distinct;ventral mesothoracic and metathoracic plates yellow ocher, more distinct. Papillae of ventral, tenth abdominal, ambulatory wart distinct. Basal plate of urogomphi

36 crescentiform, anterior and lateral margins rounded, curving ventro- caudal; mesal portion slightly concave, somewhat sculptured with a few, irregularly shaped depressions;length about two-thirds the width of plate and slightly shorter than frons. Urogomphi subconical; directed distinctly upwards and turned slightly inwards; apex sharply recurved and pointed; outer margins unequal, inner margins slightly equal; about one-third as long as basal plate; dark brown to black. Fourth instar:Average length of larva, about 18 mm. Maximum width of body, about 2.35 mm; maximum thickness of body, about 2.35 mm. Average width of head capsule, 1.46mm; range of width of head capsule, 1.24 mm to 1.64 mm. Anterior width of prothoracic shields, about 1.76 mm. Width of basal plate of urogomphi, about 1.29 mm. Setae on mesothoracic and metathoracic plates nearly lacking. Two small papillae appearing, one on each side of dorsal mesal line of first abdominal segment. Papillae on second abdominal, ambulatory wart more numerous and distinct. Fifth instar:Average length of larva, about 17 mm. Maximum width of body, about 2.68 mm; maximum thickness of body, about 2.52 mm. Average width of head capsule, 1.63mm; range of width of head capsule, 1.41 mm to 1.81 mm. Anterior width of prothoracic shields, about 1.95 mm. Width of basal plate of urogomphi, about 1.43 mm. Abdomen becoming light blue gray. Sixth instar:(Plate XII, Figures C to E; Plate XIII, Figures A to H; Plate XIV, Figures A and B; Plate XV, Figure A). Aver- age length of larva, about 16 mm. Maximum width of body, about 2.85 mm; maximum thickness of body, about 2.35 mm. Average width of head capsule, 1.87 mm; range of width of head capsule, 1.68 mm to 2.05 mm. Anterior width of prothoracic shields, about 2.27 mm. Width of basal plate of urogomphi, about 1.02mm. Epi- cranial halves of head capsule slightly rugose.First abdominal spiracle slightly larger than remaining abdominal spiracles. Abdomen, blue gray.

Dolichopodidae:Medeteraspp. Specimens of adults from the Intermountain Region were identified by Dr. R. H. Foote, United States National Museum,as mostly M. aldrichiiWh., but some were identified asM.sp. (nearnigripes Lev.) andM.sp. (nearoregonensisVan Duzee).4 Specimens from the Northwest were identified by G. Steyskal, United States Depart- ment of Agriculture, asM. aldrichiiSince the classification of this

'Personalcommunication,M. MFurniss,United States Forest Service, Ogden, Utah

37 0.25 mm

PLATE XII Temnochila virescens chlorodia Mann : A, second instar, dorsal view of thorax; B, secondinstar,dorsal view of basal plate and urogomphi; C, sixth instar, dorsal view of basal plate and urogomphi;D, sixth instar, lateral view of basal plate and urogomphi; E, sixthinstar, mesothoracic leg. C. e V u M, 02 um 1 OImm

` A, dorsal view of head; 'A.. D, abdominal spiracle; E, t C) instar: head;

sixth I I ` PLATE X I I I Mann., F, chalaza and base of abdominal seta; G, ventral IMM H, dorsal view of mandible.

7 Tom' antenna; 0.5 MM mandible: Temnochila virescens chlorodia of B, lateral view of head; C, ventral view dorsal view of view of 1 PLATE XIV Temnochila virescens chlorodiaMann.,sixth instar: A, dorsal view of com- pletelarva;B, lateral view of complete larva. group of flies is in a state of revision, no effort was made to identify the larvae down to the species level (10). Species of this genus have been reportedas being predaceous on many species of Scolytidae. mainlyIlcndroctonus, Scoh'tus,and I ps:and on Ceranlbvcidae, Ruprestidae, and Curculionidae. Records indicate thatAl. aldrichiiis widely distributed throughout the coniferous forests of western United States and British Columbia (1, i, 9). Maggots of this insect are very abundant under the bark oftrees infestedby D. pseudotsugae.It is the most numerous of all of the predators, and ranks second in number only to the parasite C.brunneri.Despite its large numbers, this species is probably notso effective as E.sphegeusin the control of the Douglas-fir beetle population. DeLeon (8, p. 73) reported that the larvae ofAl. aldrichiifind their preyfortuitously.Having no legs, the larvae are not very mobile and hence not very efficient in finding their hosts. The observed life cycle of this insect is brieflyasfollows: Adults have been observed moving very rapidlyover the hark of trees which the Douglas-fir beetle in the Intermountain Region is initially invad- ing. The emergence of the fly in the PacificNorthwestappears to be considerably laterthanthe initial attack by the bark beetle. Mating apparently takes place on the bark of infested material and theeggs are laid in small clusters under scales of thebark.Adults are present throughout thesummer.There is only one generationper year with all three instars present throughout thesummer (10). Pupation usually occurs under the bark in spring. The following descriptions were taken from the literatureon M. aldrichiiWh. by Bedard(1,pp. 40-42), DeLeon (8, pp. 62-67), and Hopping (9) with some additional observations. A drawing (Plate XV, Figure B) was made illustratinga complete, mature larva ofMedeterasp. Descriptions of immaturestages. The egg, three larval instars, and pupa are described. Egg: Average length, 0.86 nun:average width, 0.18 mm, Form suhcylindrical, with a slightly curved longitudinal axis,convex dorsally, concave ventrally, pointed atone end: margins equal: chorion smooth and shining: color at first pearly white, becoming brownish orange with development. First instar:Average length of larva, about 0.7mm. Form slender, cylindrical; creamy white in color; metapneustic; numberof body segments not determined;no sclerotized areas on the head or first thoracic segment.

41 B C

PLATE XV A, sixthinstar,ventral view of complete larva of Temnochila virescens chlor- odiaMann. ;B, mature larva, lateral view of complete larva of Medetera sp.; C, mature larva, lateral view of complete larva of Lonchaea sp Second instar: DeLeon (8, p. 62) states, "The second instar was not observed for this species. The larvae ofAl. signaticornisLoew, which were studied by the writer with greater care, went through three instars, and it is probable that there is the same number for M. aldrichii.The second instar of the former species resembles the first instar in lacking the sclerotized areas of the head and first thoracic segment." Johnsey observed a second instar for Medetera sp. associated with Dendroctonus pseudotsugae in Washington and Oregon. Again, the sclerotized areas of the head and first thoracic segment were not present(10). Third instar:(Plate XV, Figure B). Average length, 8.5 mm. Form cylindrical with a slightly curved longitudinal axis, tapering slightly at each end with the tapering more pronounced cephalad; abdomen with a rather rigid cuticle, glabrous (although DeLeon mentionsveryminute sensillae on the bodysegments),grayish white; three thoracic and eight abdominal segments; abdominal segments five to ten are subdivided at the anterior margin, forming a narrow ring on all sides except the ventral; tenth segment is about 1.4 times as long as segment nine, and about 2.5 times as long as segment 11; each of the abdominal segments, except the eighth, with a ventral pseudopod; eighth abdominal segment with the anal opening and abdominal spiracle. Head subdivided, exserted; caudal region of the dorsum of the posterior region of the head with a brownish, sclerotized, oval plate. Anterior region of head protruding forward in the form of two lateral lobes; lobes with the mandibles, mandibular plates, palpi, and other sensillae; a median spine extending anteriorly between the two lobes; internally and extending back into the mesothorax are two meta- cephalic and two tentorial black rods. Antennae apparently one- segmented, located on a pair of lateral lobes of the anterior head region. Anteriormarginof prothorax dorsally witha small crescent- shaped, sclerotized plate. Amphipneustic type of respiratory system; two pair of spiracles; cephalic pair small, located below the median lateral, longitudinal line and behind the median transverse line of the prothorax; caudal pair located in the center of the four posterior protuberances of the eighth abdominal segment,larger thanthecephalicpair, more strongly pigmented with brown, roughly oval in shape with two large oval openings into the stigmatic chamber; dorsal and lateral areas around the caudal pair of spiracles are slightly sclerotized with four branched spines located in this region close to the margin of the spiracles.

4 Eighth abdominal segment divided dorso-ventrally by a median groove and laterally by a transverse groove, thus forming four protuberances; two ventral protuberances project beyond the dorsal ones so that the former can be seen when viewed from above;also closer together and more sharply tapered than the dorsal pair of protuberances. Anus opens as a longitudinal slit on the ventral surface of, and slightly anterior to, the eighth abdominal segment, in a more or less circular padlike swelling of the cuticula. Pupa:Average length, about 4.5 mm; average width, about 2 mm; typically exarate; creamy white except for brownish sclerotized areas. Pair of sclerotized respiratory spines or horns directly behind the eyes with their bases extending parallel to the posterior margin of the eyes beneath the cuticula; pair of spiracles or vestigial spiracles on all but seventh and eighth abdominal segments. Single transverse row of spines located dorsally on all but eighth abdominal segment. Anterior to each row of dorsal abdominal spines an area of innumerable, minute, sharply pointed spines that diminish in number towards anterior margin of each segment; laterad, spines become smaller but are larger on the lateral swellings, forming distinct scabrous areas. Eighth abdominal segment with a series of five to seven pair of elongated spines; an inner, much shorter pair; all but inner and shortest pair hooked at end. Thorax with three pair of small spines; anterior spine in some instances doubled, so that on one side of pupa two spines may be present where there is normally one. Wings extend to apex of second abdominal segment, apicesof prothoracic tarsi to apex of third abdominal segment, those of mesothoracic tarsi to apex of fourth abdominal segment, those of metathoracic tarsi to apex of fifth abdominal segment. Usually pupates in a white, silky-lined pupal cell. Lonchaeidae :Lonchaeaspp. Adult specimens from the Intermountain Region were determined by Dr. C. W. Sabrosky of the United States National Museum and Dr. J. F. McAlpine of Ottawa, Canada. As in the case of Medetera,there was more than one species involved. TheLonchaea have been identified asL.sp. (nearcorticisTaylor) andL.sp. (near watsoni'Curran),'and those from the Northwest as belonging to the L. watsonigroup(10). Very little has been reported concerning this group of flies. Bedard (1, p. 49-49) included very brief descriptions and illustra-

s Personal communication, M. M Furniss. tions of the egg, mature larva, and pupa ofL. corticis.Species of this genus have been observed in association withD. pseudotsugaein Douglas-fir andD. engelmanniin Engelmann spruce in the Inter- mountain Region. It has been found under Douglas-fir bark in the Pacific Northwest. Bedard (1, p. 47) in 1933 stated, "The larvae of L.corticisunquestionably destroy much of theD pseudotsugae broods, possibly even more thanMedetera,as the larvae are more abundant in the infested trees than those of the latter species." From observations made in western Oregon, however, it appears that the larvae of this fly are more scavengers than predators. There are two generations per year and the adults of the first generation begin to emerge during the last week of June. The following descriptions were taken from the literature by Bedard(1.p. 48-49) on L.corticis,with additional information from observation. A drawing (Plate XV, Figure C) of a complete larva is included. Descriptions of immaturestages. The egg, mature larva, and pupa are described. Egg: Average length, 0.86 mm; range in length, 0.80 mm to 0.88 mm. Average width, 0.17 mm; range in width, 0.15 mm to 0.20 mm. Form subcylindrical, with a slightly curved longitudinal axis,tapering at one end, tapered end truncated; margins unequal; chorion smooth and shining; color pearly white. Mature larva:(Plate XV, Figure C). Average length, about mm. Form wedge-shaped or muscidiform, with a slightly curved longitudinal axis, tapering gradually to a sharply pointed cephalic end; creamy white and glabrous; three thoracic and eight abdominal segments; pseudopodia situated ventrally on second to eighth abdominal segments; setulae present about margins of anal area on eighth abdominal segment. Head may be retractile; with small antennae and buccal appendages; mouthparts mainly internal, visible; anterior hooks black; posterior rods extending into the prothorax and mesothorax, dark brown, fused at two locations, branching caudally. Head and prothorax without sclerotized plates. Amphipneustic type of respiratory system; two pair of spiracles; cephalic pair small, nine-lobed, located laterad and caudal on the prothorax; caudal pair located on dorso-caudal end of bluntly pointed eighth abdominal segment, containing three slits situated at right angles to each other, heavily sclerotized on dorsal portion extending outward and upward to form a heavy, blunt, spine curving above each spiracle.

45 Pupa:Average length,about5.0 mm;typicallycoarctate; puparium reddish brown, average length, about 5.2 mm; pupa creamy white, slightly shining; apices of wings extend to apex of third and last abdominal segment; apices of prothoracic tarsi to middle of same segment; apices of mesothoracic tarsi extend to apex of penultimate abdominal segment; those of metathoracic tarsi to middle of apical abdominal segment.

Braconidae:Coeloides brunneriViereck Identification of the immature stages ofCoeloides brunneriVier. was very briefly worked on by Bedard (1, p. 52). Recently, Ryan (21) published descriptions of the five larval instars. This insect has been reported as being parasitic mainly on Dendroctonus pseudotsugae (1, p. 51;22). Ryan and Rudinsky (22, p. 755) reported that a small percentage of parasitism occurs on Melanophila drummondi.It is the most abundant of all of the predators and parasites ofD. pseudotsugae.In small trees and towards the top of larger trees, C.brunnerican be very effective in parasitizing a large percentage of the Douglas-fir brood. Virtually nothing is known about the biology of this species in relation to the Douglas-fir beetle in southern Idaho, although adults have been observed on the bark of infested trees in early July. Ryan and Rudinsky (22, p. 755) reported that C.brunneriin Oregon "suc- cessfully parasitizes the Douglas-fir beetle only when the host is in the second, third, and fourth instars." There are three generations per year in western Oregon with varied incidence of diapause. The incidence of diapause increases as the season progresses, being only 5%in the first generation,45%oin the second one, and 95% in the third. Records indicate that the parasite is widely distributed from British Columbia toCalifornia and eastwardto Montana and Colorado(15, p.160). The following descriptions were taken from the literature on C. brunneriby Bedard(1, p. 52)and Ryan(21).Illustrations (Plate XVI, Figures C and D) were redrawn from Ryan. Descriptions of immaturestages. The egg, five larval instars, and pupa are described. Egg:Average length, 1.3 mm; average width at widest point, 0.15 mm. Form cylindrical with a long tapering tail; widest portion approximately one-fifth of distance from broad, rounded anterior end; tail usually slightly curved and tapering to a small acute point;

46 A

0 N

PLATE XVI A, mature larva, lateral view of complete larva of Pteromalidae; B, mature larva, frontal view of head of Pteromalidae; C, fifthinstar, frontal view of head of Coeloides brunneriVier. ; D, fifthinstar, lateral view of complete larva of C. brunneri Vier. chorion thin, elastic, translucent, without ornamentation; color white, ends becoming clear and transparent. First instar:Average length, about 1.1 mm. Form cylindrical, tapering gradually towards caudal end; distinct head and 13 body segments; first 11 body segments with a prominent band of sensory setae around the middle in addition to several smaller scattered setae; twelfth and thirteenth body segments have only a few scattered setae; body membranous and white. Head exserted; lateral margins i ndorsal view nearly straight for approximately two-thirds the length of head, converging slightly anteriorly and merging with the broadly rounded cephalic margin; antennae conical in a cephaloteral position; labrum, maxillae, and labium grouped about mouth opening as a cup-shaped, prominent, cephaloventral projection from the head capsule proper. Spiracles circular; one pair on prothorax and abdominal segments one to eight; all located slightly dorsad of midlateral line, that on thoracic segment just cephalad of caudal margin of that segment, while those on the abdominal segments lie just caudad of the cephalic margins of their respective segment. Second, third, and fourth instars:Average length of second, third, and fourth instars, 1.4, 2.3, and 3.3 mm, respectively. Bod fusiform; no prominent bands of setae present, although from four to eight individual setae aligned around center of each thoracic segment; abdominal segments with single setae present laterad of mid- dorsal and midventral lines and at the midlateral position. Head capsule spherical; outline of head in dorsal view curved with widest part in the center; mouth parts do not project. Fifth instar:(Plate XVI, Figures C and D). Average length, about 6 mm. Body fusiform to cyphosomatic; seven dorsal protrusile areas in the successive intersegmental areas, the most anterior between the metathoracic and first abdominal segment;in living larvae the first eight abdominal segments have swellings in the midlateral areas; sensory setae slightly more numerous than preceding instars; numerous cuticular spines covering the first 12 body segments except along the intersegmental lines and on the dorsal protrusile areas, also not present on terminal body segment or on the head except ventral surface. Head in frontal view obovate, constricted slightly below the midline by the hypostoma, a slight depression on either side of the dorsal midline of the epicranium; bases of antennae surrounded by a broadly elliptic antennal foramina; six pair of setae dorsad of mouth armature; labiobase with three pair of setae dorsally and many cuticular spines ventrally;lines between

48 mouthparts and parts of head capsule heavily sclerotized, stipital sclerome between cardo and stipes. Pupa:Average length, about 5.3 mm; typically exarate pupa; white in color, becoming darker with maturity; usually pupates in a white silken cocoon.

Pteromalidae:Roptrocerus eccoptogasteriRatz.,Ceci- dostiba burkeiCraw., and C.dendroctoniAsh. Adult specimens of this family which emerged from Douglas- fir slabs were determined by B. D. Burks, United States National Museum.' The parasites were identified asRoptrocerus eccoptogasteri Ratzeburg,Cecidostiba burkeiCrawford, and C.dendroctoniAsh- mead. R. eccoptogasteri has been reported as occurring in some of the Atlantic States and the Pacific Northwest. Its hostsareDendroc- tonusandIps (15, p.549). C.burkeiappears to parasitize species of Dendroctonusand occurs in the Pacific Northwest(15, p.557). Records on C.dendroctonusstate that this species is parasitic on Dendroctonus, Ips,andCylindrocopturusand occurs in West Vir- ginia and the Pacific Northwest(15, p.557). These species are not very abundant in association withD. pseudotsugae.The biology and habits of this group are presumably similar to those of C.brunneri. The following description is from characters by Michener (14, pp. 993-995), and Parker,' and from observation. Drawings (Plate XVI, Figures A and B) were made from preserved specimens.

Mature larva: (Plate XVI, Figures A and B). Body form somewhat crescent-shaped to cyphosomatic, tapering sharply towards caudal end; distinct head and 13 body segments; abdominal segments one to four wider than remaining abdominal and thoracic segments; no dorsal protrusile areas in the successive intersegmental areas; a few sensory setae located dorsally on thoracic and abdominalsegments; numerous cuticular spines scattered over the first 12 body segments; six setae around anus; body membranous and white. Head exserted, in frontal view broadly obovate, constricted slightly below the midline by the hypostoma; short, conical antennae with bases surrounded by a broadly elliptic antennal foramina; lines between mouthparts and parts of head capsule not (or feebly) sclerotized, cardo and stipes fused or nearly so, withouta conspicuous stipital Personal communication from M. M. Furniss ' Personal communication, D. E Parker, United States Forest Service, Ogden, Utah sclerome between them. Spiracles circular; one pair on mesothorax, metathorax and abdominal segments one to seven;all except first abdominal spiracle located on midlateral line, rthat of first abdominalL.I" segment slightly dorsad of midlateral line; all spiracles near cephalic margin of their respective segment.

ACKNOWLEDGMENTS This study is an outgrowth of a 1960 cooperative undertaking between Oregon State University and the Intermountain Forest and Range Experiment Station, United States Forest Service, Ogden, Utah, on the distribution of the parasites and predators of the Douglas-fir beetle infesting standing trees. Financial support for identification and description was provided by the Oregon Agricul- tural Experiment Station and National Science Foundation, grant No. G-14296. The work was clone by research assistant L. N. Kline, under the direction of J. A. Rudinskv. Specimens from the Inter- mountain Region were collected from the Boise National Forest and layette National Forest in Idaho, while L. N. ]\-line was employed by the United States Forest Service, and Pacific Northwest specimens were taken from the Marys Peak watershed near Corvallis, Oregon. The authors acknowledge the assistance of D. H. Parker, United States Forest Service, Ogden, Utah, who originated the distribution project; M. M. Furniss, of the same agency, who helped in planning and in the field; and P. O. Ritcher, head of the Department of Ento-.7, mology, Oregon State University, who checked many of the descriptions.

REFERENCES CITED 1. Bedard, W. D.,The Douglas-fir beetle, its seasonal history, biology, habits, and control.USDA, Forest Service, Forest Insect Field Station, Coeur d' Alene, Idaho, 1933 (Processed.) 2. Blackwelder, Richard E, Fourth supplement to theLeng Catalogue of Coleoptera of America, North of MexicoMount Vernon, N. Y.: Frey- bourg, 1939. 3. Blackwelder, Richard E., and Ruth M. Blackwelder, Fifth supplement to theLeng Catalogue of Coleoptera of America, North of Mexico.Burling- ton, Vt.: Lane Press, 1948 4. Roving, A G., The larvae ofEnoclerus leconteiWolcott andCallimerus arcuferChapin, of the beetle family Cleridae. Proc. of the Ent. Soc. of Wash., 30:94-100, 1928. 5. Roving, A. G., and A. B. Champlain, Larvae of North American beetles of the family Cleridae. Proc U. S. Natl Museum, 57:575-649, 1920.

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21. Ryan, Roger B, Durations of the immature stadia of Coeloides brunneri (Hymenoptera: Braconidae) at various constant temperatures, with descriptions of the five larval instars. Annals of the Ent. Soc. of Amer., 55:403- 409, 1962.

51 22. Ryan, Roger B., and Julius A. Rudinsky, Biology and habits of the Douglas- fir beetle parasite,Coeloides brunneriViereck (Hymenoptera: Braconidae), in western Oregon, The Canadian Entomologist, 94:748-763, 1962. 23 Struble, George R., The biology of certain Coleoptera associated with bark beetles in western yellow pine, Univ. of Calif. Publications in Ent. 5(6) 105-134, 1930. 24. Struble, George R, Biology of two native coleopterous predators of the mountain pine beetle in sugar pine, The Pan-Pacific Entomologist,18:97- 107, 1942. 25. Struble, George R, Laboratory propagation of two predators of the mountain pine beetle. Jour. of Econ. Ent, 35:841-844, 1942. 26. Struble, George R., and L. H. Carpelan, External sex characters of two important native predators of the mountain pine beetle in sugar pine. The Pan-Pacific Entomologist, 17:153-156, 1941. 27. Torre-Bueno, J.R., De La, ed.,A Glossary of Entomology.Brooklyn, N. Y.: Brooklyn Entomological Society, 1950. 28. Wolcott, Albert B., Catalogue of North American beetles of the family Cleridae. Fieldiana : Zoology, 32:61-105. 1947.

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